Part 24

If we compare the productions of different islands, we meet with peculiarities which throw much light on the subject of distribution. In the Galapagos islands, between 500 and 600 miles from the west coast of South America, there are thirty-two species of land-birds, all but two or three being peculiar to the group. In Madeira, about 400 miles from the coast of Morocco, there are nearly twice as many land-birds as in the Galapagos, but only two of these are peculiar to the island, the rest being South European or N. African species. The Azores are 1,000 miles west of Portugal, and they contain twenty-two species of land-birds, every one of which is European except one bullfinch which is slightly different and forms a peculiar species. This remarkable difference in the proportion of peculiar species between the Galapagos and the Atlantic islands, is well explained by the theory that land-birds rarely fly directly out to sea, except when carried against their will by storms and gales of wind. Now the Azores are situated in an especially stormy zone, and it is an observed fact that after every severe gale of wind some new bird or insect is seen on the islands. The Galapagos, on the contrary, are in a very calm sea where violent storms are almost unknown, and thus new birds from the mainland very rarely visit these islands. Madeira is less stormy than the Azores, but its comparative nearness makes up for this difference in the case of birds. In insects, however, the species of Madeira are much more peculiar (and more numerous) than those of the more distant Azores; while those of the Galapagos are few, but all peculiar, and belonging to groups many of which are widely spread over the globe. All these facts are entirely in accordance with the view that oceanic islands have been peopled from the nearest continents by various accidental causes; while they are entirely opposed to the theory that such islands are remnants of old continents and have preserved some portion of their inhabitants.

It is a curious fact, that land reptiles, such as snakes and lizards, are found in many islands where there are no mammalia or frogs; and we therefore conclude that there must be some means by which their ova can be safely carried across great widths of sea. A single peculiar frog inhabits New Zealand, and some species are found in the Pacific islands as far eastward as the Fijis, but they are absent from all other oceanic islands. Snakes also extend to the Fijis, and there are two species in the Galapagos, but none in the other oceanic islands. Lizards, however, are found in Mauritius and Bourbon; in New Zealand; in all the Pacific islands, and in the Galapagos. It is clear then that next to Mammals, frogs and toads are most completely shut out by an ocean barrier; then follow snakes, but as these are only found in the Galapagos and are very like South American species, they may possibly have been conveyed in boats or by floating trees. Lizards, however, are so wide-spread over almost all the warmer islands of the great oceans, that they must have some natural way of passing over, but the exact mode in which this is effected has not yet been discovered. Birds, as we have seen, are liable to be carried by winds and storms over great widths of sea, but this only applies to certain groups; and large numbers which feed on the ground or which inhabit the depths of the forests, are almost as strictly confined to their respective countries by even a narrow arm of the sea as are the majority of the mammalia.

This sketch of the mode in which the various kinds of islands have been stocked with their animal inhabitants forms the best introduction to the study of those changes in our continents which have led to the existing distribution of animals. It demonstrates the importance of the sea as a _barrier_ to the spread of all the higher animals; and we are thus naturally led on to inquire, how far and to what extent such barriers have in past time existed between lands which are now united, and on the other hand what existing oceanic barriers are of comparatively recent origin. In pursuing this inquiry we shall have to take account of those grand views of the course of nature associated with the names of Lyell and Darwin--of the slow but never-ceasing changes in the physical conditions, the outlines and the mutual relations of the land-surfaces of the globe; and of the equally slow and equally unceasing changes in the forms and structures of all organisms, to a great extent correlated with, and perhaps dependent on, the former set of changes. Combining these two great principles with other ascertained causes of distribution, we shall be enabled to deal adequately with the problem before us, and give a rational, though often only an approximative and conjectural, solution of the many strange anomalies we meet with in studying the distribution of living things.

_Past and Present Distribution of Land and Sea._--Before proceeding to give details as to the distribution of animals, it is necessary to point out certain geographical features which have had great influence in bringing about the existing state of things.

The extreme inequality with which land and water is distributed has often been remarked, but what is less frequently noted is the singular way in which all the great masses of land are linked together. Notwithstanding the small proportion of land to water, the vast difference in the quantity of land in the northern and southern hemispheres, and the apparently hap-hazard manner in which it is spread over the globe, we yet find that no important area is completely isolated from the rest. We may even travel from the extreme north of Asia to the three great southern promontories--Cape Horn, the Cape of Good Hope, and Tasmania--without ever going out of sight of land; and, if we examine a terrestrial globe, we find that the continents in their totality may be likened to a huge creeping plant, whose roots are at or around the North Pole, whose matted stems and branches cover a large part of the northern hemisphere, while it sends out in three directions great offshoots towards the South Pole. This singular arrangement of the land surface into what is practically one huge mass with diverging arms, offers great facilities for the transmission of the varied forms of animal life over the whole earth, and is no doubt one of the chief causes of the essential unity of type which everywhere characterises the existing animal and vegetable productions of the globe.

There is, moreover, good reason to believe that the general features of this arrangement are of vast antiquity; and that throughout much of the Tertiary period, at all events, the relative positions of our continents and oceans have remained the same, although they have certainly undergone some changes in their extent, and in the degree of their connection with each other. This is proved by two kinds of evidence. In the first place, it is now ascertained by actual measurement that the depths of the great oceans are so vast over wide areas, while the highest elevations of the land are limited to comparatively narrow ridges, that the mass of land (above the sea-level) is not more than ¹⁄₃₆th part of the mass of the ocean. Now we have reason to believe that subsidence and elevation bear some kind of proportion to each other, whence it follows that although several mountain ranges have risen to great heights during the Tertiary period, this amount of elevation bears no proportion to the amount of subsidence required to have changed any considerable area of what was once land into such profound depths as those of the Atlantic or Pacific Oceans. In the second place, we find over a considerable area of all the great continents fresh-water deposits containing the remains of land animals and plants; which deposits must have been formed in lakes or estuaries, and which therefore, speaking generally, imply the existence in their immediate vicinity of land areas comparable to those which still exist. The Miocene deposits of Central and Western Europe, of Greece, of India, and of China, as well as those of various parts of North America, strikingly prove this; while the Eocene deposits of London and Paris, of Belgium, and of various parts of North and South America, though often marine, yet by their abundant remains of land-animals and plants, equally indicate the vicinity of extensive continents. For our purpose it is not necessary to go further back than this, but there is much evidence to show that throughout the Secondary, and even some portion of the Palæozoic periods, the land-areas coincided to a considerable extent with our existing continents. Professor Ramsay has shown[64] that not only the Wealden formation, and considerable portions of the Upper and Lower Oolite, but also much of the Trias, and the larger part of the Permian, Carboniferous, and Old Red Sandstone formations, were almost certainly deposited either in lakes, inland seas, or extensive estuaries. This would prove that, throughout the whole of the vast epochs extending back to the time of the Devonian formation, our present continents have been substantially in existence, subject, no doubt, to vast fluctuations by extension or contraction, and by various degrees of union or separation, but never so completely submerged as to be replaced by oceans comparable in depth with our Atlantic or Pacific.

[64] _Nature_, 1873, p. 333; _Quarterly Journal of the Geological Society_, 1871, pp. 189 and 241.

This general conclusion is of great importance in the study of the geographical distribution of animals, because it bids us avoid the too hasty assumption that the countless anomalies we meet with are to be explained by great changes in the distribution of land and sea, and leads us to rely more on the inherent powers of dispersal which all organisms possess, and on the union or disruption, extension or diminution, of existing lands--but always in such directions and to such a limited extent as not to involve the elevation of what are now the profoundest depths of the great oceans.

_Zoological Regions._--We will now proceed to sketch out the zoological features of the six great biological regions; and will afterwards discuss their probable changes during the more recent geographical periods, in accordance with the principles here laid down.

_The Palæarctic Region._

The Palæarctic, or North Temperate region of the Old World, is not only by far the most extensive of the zoological regions, but is the one which agrees least with our ordinary geographical divisions. It includes the whole of Europe, by far the largest part of Asia, and a considerable tract of North Africa; yet over the whole of this vast area there prevails a unity of the forms of animal life which renders any primary subdivision of it impossible, and even secondary divisions difficult. But besides being the largest of the great zoological regions, there are good reasons for believing this to represent the most ancient, and therefore the most important centre of the development of the higher forms of animal life,--and it is therefore well to consider it first in order.

In enumerating the most important animal groups characteristic of this and other regions, it must be clearly understood that such groups are not always absolutely confined to one region. Here and there they will often overlap the boundaries, while in other cases single species may have a wide distribution in one or more of the adjacent regions; but this does not at all affect the main fact, that the group as a whole is very abundant and very widely spread over the region in question, while it is very rare, or confined to a very limited area in adjacent regions, and is therefore specially characteristic of the one as compared with other parts of the world. Bearing this in mind, we shall find that the Palæarctic region is well characterized by a considerable number of typical groups, although, as we shall presently see, it has in recent geological times lost much of its ancient richness and variety of animal life.

Among Mammalia the groups most characteristic of this region are the moles (Talpidæ), a family consisting of eight distinct genera which range over the whole region, but beyond it barely enter the Oriental region in North India, and the Nearctic region in North-West America; camels, confined to the deserts of North Africa and Asia; sheep and goats (Capra), only found beyond the region in the Nilgherries and Rocky Mountains; several groups of antelopes, and many peculiar forms of deer; hamsters (Cricetus), sand rats (Psammomys), mole rats (Spalax), and pikas (Lagomys), with several other forms of rodents. Wolves, foxes, and bears, are also very characteristic, though by no means confined to the region.

Among birds the most important group is certainly the small-sized, but highly-organized warblers (Sylviidæ), which, although almost universally distributed, are more numerous, and have more peculiar and characteristic genera here than in any other region. Most of our song-birds, and many of the commonest tenants of our fields, woods, and gardens, belong to this family; and identical or representative species are often found ranging from Spain to China, and from Ireland to Japan. The reedlings (Panuridæ), the tits (Paridæ), and the magpies (Pica), are also very characteristic; while among the finches (Fringillidæ), a considerable number of genera are peculiar. A large number of peculiar groups of grouse (Tetraonidæ), and pheasants (Phasianidæ) are also characteristic of this region. Although the reptiles and fresh-water fishes are comparatively few, yet many of them are peculiar. Thus, no less than 2 genera of snakes, 7 of lizards, and 16 of batrachia, are confined to the Palæarctic region, as well as 20 genera of fresh-water fishes.

The insects and land-shells offer their full proportion of peculiar types; but it would lead us beyond our special object to enter into details with regard to these less known groups of animals.[65]

[65] Details will be found in the Author’s work on _The Geographical Distribution of Animals_.

_The Ethiopian Region._

The Ethiopian region, consisting of Africa south of the Tropic of Cancer with Madagascar, is of very small area compared with the Palæarctic region; yet owing to the absence of extreme climates, and the tropical luxuriance of a considerable portion of its surface, it supports a greater number and variety of large animals than any other part of the globe of equal extent. Much of the speciality of the region is, however, due to the rich and isolated fauna of Madagascar, the peculiarities of which may be set aside till we come to discuss the past history of the Ethiopian region.

Considering then, first, the zoological features of tropical and southern Africa alone, we find a number of very peculiar forms of mammalia. Such are the golden moles, the Potamogale, and the elephant-shrews among Insectivora; the hippopotami and the giraffes, among Ungulata; the hyæna-like Proteles (Aard-wolf), and Lycaon (hyæna-dog), among Carnivora; and the Aard-varks (Orycteropus) among Edentata. These are all peculiar; but among highly characteristic forms are the baboons, and several genera of monkeys and apes; several peculiar Lemurs; a great variety of the civet-family (Viverridæ), and of rodents; peculiar genera of swine (Potamochærus and Phacochærus), and a greater abundance and variety of antelopes than are to be found in all the other regions combined. But the Ethiopian region is strikingly distinguished from all others, not only by possessing many peculiar forms, but by the absence of a number of common and widely distributed groups of mammalia. Such are--the bears, which range over the whole northern hemisphere, and as far south as Sumatra in the eastern and Chili in the western hemisphere, yet they are totally wanting in Tropical and South Africa;--the deer, which are still more widely distributed, ranging all over North and South America, and over all Asia to Celebes and the Moluccas, yet they are totally absent from the Ethiopian region; goats and sheep, true oxen (Bos), and true pigs (Sus), are also absent; though as to the last there is some doubt, certain wild pigs having been observed, though rarely, in various parts of Tropical Africa, but it is not yet determined whether they are indigenous, or escaped from domestication. The absence of such wide-spread families as the bears and deer is, however, most important, and must be taken into account when we come to consider the geographical changes needed to explain the actual state of the Ethiopian fauna.

The birds are not proportionately so peculiar, yet there are many remarkable forms. Most important are the plantain-eaters, the ground-hornbills, the colies, and the anomalous secretary-bird;--while among characteristic families there are numbers of peculiar genera of flycatchers, shrikes, crows, sun-birds, weaver-birds, starlings, larks, francolins, and the remarkable subfamily of the Guinea-fowls. There are not such striking deficiencies among birds as among mammals, yet there are some of importance. Thus, there are no wrens, creepers, or nut-hatches, and none of the wide-spread group comprising the true pheasants and jungle fowl--a deficiency almost comparable with that of the bears or the deer. Among the lower vertebrates there are 3 peculiar families of snakes and 1 of lizards, as well as 1 of toads and 3 of fresh-water fishes.

_The Oriental Region._

The Oriental region comprises all tropical Asia east of the Indus, with the Malay Islands as far as Java, Borneo, and the Philippines. In its actual land-area it is the smallest region except the Australian; but if we take into account the wide extent of shallow sea connecting Indo-China with the Malay Islands, and which has, doubtless, at no distant epoch, formed an extension of the Asiatic Continent, it will not be much smaller than the Ethiopian region. Here we find all the conditions favourable to the development of a rich and varied fauna. The land is broken up into great peninsulas and extensive islands; lofty mountains and large rivers everywhere intersect it; while along its northern boundary stretches the highest mountain-range upon the globe. Much of this region lies within the equatorial belt, where the equability of temperature and abundance of moisture produce a tropical vegetation of unsurpassed luxuriance. We find here, as might be expected, that the variety and beauty of the birds and insects is somewhat greater than in the Ethiopian region; although, as regards mammalia, the latter is the most prolific, both in genera, species, and individuals.

The families of Mammalia actually peculiar to this region are few in number, and of limited extent. They are,--the Galeopithecidæ, or flying lemurs; the Tarsiidæ, consisting of the curious little tarsier, allied to the lemurs; and the Tupaiidæ, a remarkable group of squirrel-like Insectivora. There are, however, a considerable number of peculiar genera, forming highly characteristic groups of animals--such as the various apes, monkeys, and lemurs, almost all the genera of which are peculiar; a large number of civets and weasels; the beautiful deer-like Chevrotains, often called mouse-deer; and a few peculiar antelopes and rodents. It must be remarked that we find here none of those deficiencies of wide-spread families which were so conspicuous a feature of the Ethiopian region--the only one worth notice being the dormice (Myoxidæ), a small family spread over the Palæarctic and Ethiopian regions, but not found in the Oriental.

The birds of the Oriental region are exceedingly numerous and varied, there being representatives of about 350 genera of land-birds, of which nearly half are peculiar. Three families are confined to the region--the hill-tits (Liotrichidæ), the green bulbuls (Phyllornithidæ), and the gapers (Eurylæmidæ); while four other families are more abundant here than elsewhere, and are so widely distributed throughout the region as to be especially characteristic of it. These are--the elegant pittas, or ground-thrushes (Pittidæ), the trogons (Trogonidæ), the hornbills (Bucerotidæ), and the pheasants (Phasianidæ); represented by such magnificent birds as the fire-backed pheasants, the ocellated pheasants, the Argus-pheasant, the pea-fowl, and the jungle-fowl.

Reptiles are very abundant, but only 3 small families of snakes are peculiar. There are also 3 peculiar families of fresh-water fishes.

_Past Changes of the Great Eastern Continent._--Having thus briefly sketched the main features of the existing faunas of Europe, Asia, and Africa, it will be well, while their resemblances and differences are fresh in our memory, to consider what evidence we have of the changes which may have resulted in their present condition. All these countries are so intimately connected, that their past history is greatly elucidated by the knowledge we possess of the tertiary fauna of Europe and India; and we shall find that when we once obtain clear ideas of their mutual relations, we shall be in a better position to study the history of the remaining continents.

Let us therefore go back to the Miocene or middle tertiary epoch, and see what was then the distribution of the higher animals in these countries. Extensive deposits, rich in animal remains of the Miocene age, occur in France, Switzerland, Germany, Hungary, Greece; and also in North-Western India at the Siwalik Hills, in Central India in the Nerbudda Valley, in Burmah, and in North China; and over the whole of this immense area we find a general agreement in the fossil mammalia, indicating that this great continent was probably then, as now, one continuous land. The next important geographical fact that meets us, is, that many of the largest and most characteristic animals, now confined to the tropics of the Oriental and Ethiopian regions, were then abundant over much of the Palæarctic region. Elephants, rhinoceroses, tapirs, horses, giraffes, antelopes, hyænas, lions, as well as numerous apes and monkeys, ranged all over Central Europe, and were often represented by a greater variety of species than exist now. Antelopes were abundant in Greece, and several of these appear to have been the ancestors of those now living in Africa; while two species of giraffes also inhabited Greece and North-West India. Equally suggestive is the occurrence in Europe of such birds as trogons and jungle-fowl characteristic of tropical Asia, along with parrots and plantain-eaters allied to forms now living in West Africa.

Let us now inquire what information Geology affords us of changes in land and sea at this period. From the prevalence of early tertiary deposits over the Sahara and over parts of Arabia, Persia, and Northern India, geologists are of opinion that a continuous sea or strait extended from the Bay of Bengal to the Atlantic Ocean, thus cutting off the Peninsula of India with Ceylon, as well as all tropical and South Africa from the great northern continent.[66] At the same time, and down to a comparatively recent period, it is almost certain that Northern Africa was united to Spain and to Italy, while Asia Minor was united to Greece, thus reducing the Mediterranean to the condition of two inland seas. We also know that the north-western Himalayas and some of the high lands of Central Asia were at such a moderate elevation as to enjoy a climate as mild as that which prevailed in Central Europe during the Miocene epoch,[67] and was therefore perhaps equally productive in animal and vegetable life.

[66] Mr. Searles V. Wood, “On the Form and Distribution of the Land-tracts during the Secondary and Tertiary Periods respectively,” _Philosophical Magazine_, 1862.